REVIEW

J Anim Sci Technol 2023;65(1):16-31 Journal of Animal Science and Technology

https://doi.org/10.5187/jast.2022.e114 pISSN 2672-0191 eISSN 2055-0391

The roles of growth factors and

hormones in the regulation of
muscle satellite cells for cultured
meat production
Syed Sayeed Ahmad1,2#, Hee Jin Chun1#, Khurshid Ahmad1,2,
Sibhghatulla Shaikh1,2, Jeong Ho Lim1,2, Shahid Ali1,2, Sung Soo Han2,3,
Sun Jin Hur4, Jung Hoon Sohn5, Eun Ju Lee1,2* and Inho Choi1,2*
1
Department of Medical Biotechnology, Yeungnam University, Gyeongsan 38541, Korea
2
Research Institute of Cell Culture, Yeungnam University, Gyeongsan 38541, Korea
3
School of Chemical Engineering, Yeungnam University, Gyeongsan 38541, Korea
4
Department of Animal Science and Technology, Chung-Ang University , Anseong 17546, Korea
5
Synthetic Biology and Bioengineering Research Center, Korea Research Institute of Bioscience and
Biotechnology (KRIBB), Daejeon 34141, Korea

Received: Oct 31, 2022 Abstract

Revised: Nov 25, 2022
Accepted: Nov 27, 2022
Cultured meat is a potential sustainable food generated by the in vitro myogenesis of mus-
cle satellite (stem) cells (MSCs). The self-renewal and differentiation properties of MSCs
#These authors contributed equally to
this work. are of primary interest for cultured meat production. MSC proliferation and differentiation are
influenced by a variety of growth factors such as insulin-like growth factors (IGF-1 and IGF-
*Corresponding author
Eun Ju Lee 2), transforming growth factor beta (TGF-β), fibroblast growth factors (FGF-2 and FGF-21),
Department of Medical Biotechnology, platelet-derived growth factor (PDGF) and hepatocyte growth factor (HGF) and by hormones
Yeungnam University, Gyeongsan
38541, Korea. like insulin, testosterone, glucocorticoids, and thyroid hormones. In this review, we investi-
Tel: +82-53-810-3589 gated the roles of growth factors and hormones during cultured meat production because
E-mail: gorapadoc0315@hanmail.net
these factors provide signals for MSC growth and structural stability. The aim of this article is
Inho Choi
Department of Medical Biotechnology,
to provide the important idea about different growth factors such as FGF (enhance the cell
Yeungnam University, Gyeongsan proliferation and differentiation), IGF-1 (increase the number of myoblasts), PDGF (myoblast
38541, Korea.
Tel: +82-53-810-3024 proliferation), TGF-β1 (muscle repair) and hormones such as insulin (cell survival and growth),
E-mail: inhochoi@ynu.ac.kr testosterone (muscle fiber size), dexamethasone (myoblast proliferation and differentiation),
Copyright © 2023 Korean Society of and thyroid hormones (amount and diameter of muscle fibers and determine the usual pattern
Animal Sciences and Technology. of fiber distributions) as media components during myogenesis for cultured meat production.
This is an Open Access article
distributed under the terms of the Keywords: Muscle satellite cells, Growth factors, Hormones, Myogenesis, Cultured meat
Creative Commons Attribution
Non-Commercial License (http://
creativecommons.org/licenses/by-
nc/4.0/) which permits unrestricted INTRODUCTION
non-commercial use, distribution, and
reproduction in any medium, provided In vitro cultured meat production is an innovative concept based on tissue engineering and myogenesis
the original work is properly cited.
without slaughtering of livestock [1–3]. These meats are produced from muscle satellite (stem) cells
ORCID
Syed Sayeed Ahmad
(MSCs), which are being widely used for this purpose because they proliferate and differentiate
https://orcid.org/0000-0002-2829-2768 efficiently and are primary generators of new myonuclei [4]. MSCs are important for the maintenance

16 https://www.ejast.org

Hee Jin Chun
https://orcid.org/0000-0002-4171-7062
Khurshid Ahmad
https://orcid.org/0000-0002-1095-8445
Sibhghatulla Shaikh
https://orcid.org/0000-0002-7489-2393
Jeong Ho Lim
https://orcid.org/0000-0002-0375-8170
Shahid Ali
https://orcid.org/0000-0002-4724-5086
Sung Soo Han
https://orcid.org/0000-0003-0773-2661
Sun Jin Hur
https://orcid.org/0000-0001-9386-5852
Jung Hoon Sohn
https://orcid.org/0000-0002-5311-6424
Eun Ju Lee
https://orcid.org/0000-0003-2496-0463
Inho Choi
https://orcid.org/0000-0002-0884-5994
Competing interests
No potential conflict of interest relevant to
this article was reported.
Funding sources
This work was supported by the Korea
Institute of Planning and Evaluation for
Technology in Food, Agriculture, Forestry
(IPET) through High Value-added Food
Technology Development Program funded
by Ministry of Agriculture, Food and Rural
Affairs (MAFRA)(321026-05 and 322008-
5). This research was also supported by the
Basic Science Research Program through
the National Research Foundation of Korea
(NRF) funded by the Ministry of Education
(2020R1A6A1A03044512).
Acknowledgements
Not applicable.
Availability of data and material
Upon reasonable request, the datasets
of this study can be available from the
corresponding author.
Authors’ contributions
Conceptualization: Ahmad SS, Chun HJ.
Data curation: Ahmad K, Shaikh S, Lim JH,
Ali S.
Formal analysis: Ahmad K, Shaikh S, Lim
JH, Ali S, Han SS, Hur SJ, Sohn JH, Lee
EJ, Choi I.
Writing - original draft: Lee EJ, Choi I.
Writing - review & editing: Ahmad SS, Chun
HJ, Ahmad K, Shaikh S, Lim JH, Ali S,
Han SS, Hur SJ, Sohn JH, Lee EJ, Choi I.
Ethics approval and consent to participate
This article does not require IRB/IACUC
approval because there are no human and
animal participants.
https://doi.org/10.5187/jast.2022.e114
Ahmad et al.
of skeletal muscle (SM) structure and function, and meat is largely composed of SM. The SM
contains 90% muscle fibers and 10% connective and fat tissues [5–8]. Furthermore, meat produced
using MSCs represents an excellent new source of high-quality protein [9–11]. MSC self-renewal
capacity is important for maintaining MSC populations and generating enormous numbers of
myogenic cells, which proliferate, divide, fuse, and contribute to the synthesis of new myofibers [12].
Cultured meat was initially produced using bovine MSCs [13], and considerable research has been
undertaken in recent years to produce functional meat products [14].
Myogenesis is a term used for the process leading from the development to the formation
of SM tissue and involves MSC activation and proliferation (due to the expressions of muscle-
specific genes and cell cycle-related factors) and the fusion of differentiating myoblasts into mature
myofibers (under the direction of muscle regulatory factors). Myogenesis is mediated by MSCs
and is responsible for the normal growth and repair of SM. This process is regulated by myogenic
regulatory factors such as Pax3, Pax7, Myf5, MyoD, MyoG, and Myf6 [15,16] (Fig. 1).
Furthermore, MSC culture [17,18] and the directed differentiation of pluripotent MSC [19,20]
provide novel approaches to in vitro myogenesis. Several extracellular matrix (ECM) proteins such
as fibromodulin (FMOD) [7,21], matrix gla protein [22], and dermatopontin (DPT) [23] have
been reported to regulate myogenesis, and ECM contains molecules such as collagen, integrin,
decorin, biglycan, DPT, FMOD, fibronectin, glycosaminoglycan, laminin, and dystrophin that
provide structural support, cellular communication, and contribute much to the architectural
maintenance of SM [5,24].
Cultured meat is an attractive option compared with regular meat as it is free from environmental
pollution, ethical issues, and food security challenges [25–27]. However, several issues, such as color,
texture, flavor, and nutritional value, which all depend on protein and intramuscular fat contents,
remain to be resolved [28,29]. Cattle, chickens, pigs, and fish have been reported to be suitable
target species for biopsy leading to cultured meat production [25], and it has been established that a
single biopsy may replace the killing of 20 animals [30].
The different types of MSC, namely, adult, pluripotent (PSCs; embryonic MSC), and induced
pluripotent MSC) are used to produce cultured meat [31]. MSCs and adipocytes are required for
initiating cultured meat production and, in combination, may contribute to achieving an original
meat flavor and texture [12,32,33]. Porcine SM multipotent progenitor cells have a greater doubling
capacity than MSCs, but they require expensive recombinant growth factors (GFs) and do not
develop into SM fibers as effectively as MSCs [34]. Accordingly, MSCs are the most common
source used for cultured meat production, and PSCs (embryonic stem cell [ESC] and iPSC) are
considered a second option.
Technical difficulties related to culture media, such as GF, hormones, identification of animals
(age and health), sampling of MSCs, consumer acceptance, and religious standpoints, pose
considerable challenges to the establishment of viable large-scale in vitro meat production systems,
and thus, more research is required before this meat culturing system can be implemented at
industrial levels.
MSC proliferation and differentiation are influenced by many GFs such as fibroblast growth
factor (FGF), insulin-like growth factors (IGF-1 and 2), transforming growth factor-beta (TGF-
β) [35], platelet-derived growth factor (PDGF), and hepatocyte growth factor (HGF). GFs can
manage growth signal responses during development and are highly active during cell proliferation
and differentiation [16,36–38]. The roles of several GFs and hormones during myogenesis for
cultured meat production are summarized in Table 1. Three key hormone types related to the
physiological system, that is, testosterone (promotes cellular growth and repair), growth hormone,
and glucocorticoids, participate in cultured meat production [39]. The roles of GFs and expressions
https://www.ejast.org | 17
Growth factors and hormones in cultured meat production

Fig. 1. Cultured meat production based on myogenesis. MSC isolated from selected animals through biopsy
or from animal tissues have different proliferation and differentiation properties. The roles of different growth and
muscle regulatory factors during myogenesis are also included in the figure. FGF2, HGF, and IGF-1 enhance
cell proliferation, and FGF21 aids myotube formation. FGF, fibroblast growth factor; HGF, hepatocyte growth
factor; IGF, insulin-like growth factors; MSC, muscle satellite (stem) cell.

of muscle regulatory factors at different myogenesis stages are shown in Fig. 1, and a schematic of
the process from MSC isolation to muscle fiber development is provided in Fig. 1.
Types of GFs and hormones in culture medium critically maintain cell proliferation and
differentiation, and it has been demonstrated that in their absence, in vitro cultured MSCs lose their
stemness [4]. Thus, understanding the properties of MSCs and the effects of GFs and hormones
that control myogenesis is critical for the mass production of cultured meat. Previous studies have
shown that GFs and hormones are involved in myogenesis, and thus, in this review, we focus on
how GFs and hormones exert their influences throughout the stages of myogenesis. The objective
of this article was to summarize the roles of different GFs and hormones during MSC proliferation
and differentiation for in vitro cultured meat production.

ROLES OF GROWTH FACTORS IN MYOGENESIS FOR

CULTURED MEAT PRODUCTION
Fibroblast growth factors
FGFs are peptide hormones that belong to a large family of polypeptides locally present in
developing tissues [75] and are secreted by liver, adipocytes, pancreas, and SM [76]. The FGF

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 Ahmad et al.

Table 1. Growth factors, hormones, and other supplements and their functions during myogenesis and cultured meat production. Several GFs (IGFs,
insulin, TGF-β, FGF, PDGF, and HGF) and hormones (thyroid hormones, testosterone, and glucocorticoids) and other supplements (heme proteins,
transferrin, BMP, and mTOR) and their functions during cultured meat production and myogenesis are mentioned
Myogenic
Embryonic Tissue
Supplements Function progenitor References
MSC engineering
cells
IGF-1 · Induces the proliferation and differentiation of cultured myoblast and V V V [40–44]
promotes tissue growth and development
IGF-2 · Stimulate both proliferation and differentiation of myoblasts and V V [42,45]
MSC.
TGF-β · Helps in collagen type I synthesis in the ECM, and supporting myofi- V V [46–48]
ber development
FGF2 · Activate myogenic progenitor cells and promotes proliferation and V V V [49–52]
growth of mouse myoblasts.
· Upregulates Myf5 and MyoD.
· Increases the MSC proliferation.
FGF21 · Involved in cell proliferation and differentiation. V [53,54]
Insulin · Promotes MSC survival, self-renewal, and myoblast growth. V V [43,55–58]
· Regulators of muscle protein synthesis and hypertrophy.
HGF · Activation of quiescent MSC in vivo upon muscle injury. V V [52,59,60]
Testosterone · Increase in muscle fiber size and MSC number. [61–63]
Transferrin · Stimulate myogenesis and terminal differentiation in fast chicken V V V [64,65]
muscle during embryonic development.
Heme protein · Provides typical color of meat and slightly metallic taste of beef. [66,67]
(Hemoglobin and · Increase the iron content in the product.
myoglobin)
BMP · Preserve the progenitor pool by inhibiting the Myf5 and MyoD expres- [68]
sion.
mTOR · Regulate MSC activity and myogenesis by upregulating the expres- [69,70]
sion of Pax7, Myf5, MyoD, and myogenin.
Creatine · Reduces muscle damage by decreasing the inflammatory response [71,72]
and oxidative stress, and activating MSC. Helps in myopathies.
PDGF · Increases and improve MSC proliferation. V V [68,73,74]
Thyroxine · Helps to myotube in maintaining their differentiated state for a longer V V [57]
period.
GF, growth factor; IGF, insulin-like growth factors; TGF, transforming growth factor; FGF, fibroblast growth factor; PDGF, platelet-derived growth factors; HGF, hepatocyte growth fac-
tor; BMP, bone morphogenetic protein; mTOR, mechanistic target of rapamycin; MSC, muscle satellite (stem) cell; ECM, extracellular matrix.

family is composed of 22 ligands that interact with four different FGF receptors, and FGF/
FGFR signaling governs cell survival, proliferation, migration, differentiation, and tissue repair/
regeneration [77]. FGF-21 is actively involved in cell proliferation and differentiation [53]. On the
other hand, FGF-2 (basic FGF) is secreted by fibroblasts and has been used extensively for in vitro
MSC culture. FGF-2 supplementation increased mouse myoblast growth by 25%, but the removal
of FGF-2 from culture media increased the differentiation of myoblast cells. Also, the endogenous
secretion of FGF-1 (acidic FGF) by MSCs may trigger cell proliferation and muscle regeneration
[78,79]. On the other hand, FGF enhances cell proliferation and differentiation, which play
important roles during the initial stage of myogenesis. In addition, FGF-2 also promotes tissue
formation and repair by interacting with heparin sulfate cofactor [80].

Platelet-derived growth factor

PDGFs increase the production of three SM basement membrane components, namely, laminin
(70%), fibronectin (30%), and type IV collagen (70%). PDGF has four known isoforms (PDGF
A, B, C, and D), which are stabilized by intermolecular disulfide bonds. All four PDGF isoforms
contain a highly conserved 100 amino acid GF domain [81]. PDGFs are powerful mitogens and

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Growth factors and hormones in cultured meat production

affect a wide range of cells, including fibroblasts, smooth muscle cells, connective tissue, bone, and
cartilage cells, and have been demonstrated to boost mouse myoblast proliferation. Thus, PDGFs
are important components during the production of cell-cultured meat [82].

Insulin-like growth factors

IGFs play important roles in myogenesis and muscle regeneration by activating MSC proliferation,
increasing protein synthesis, and promoting differentiation [83-86]. There are two IGF types, that
is, IGF-1 and IGF-2. IGF-1 is a 70 aa long polypeptide that shares ~60% similarity with IGF-2
[87]. IGF-1 was reported to be actively involved in the proliferation and differentiation of myoblast
cells [83,88]. The negative effect of myostatin (MSTN) was found to be compensated by the
positive effect of IGF-1 during myogenesis [85]. IGF-1 promotes tissue growth and development,
stimulates cell proliferation, and has anti-aging, anti-inflammatory, and antioxidant properties [41].
IGF-1 also has direct and indirect glucose-lowering effects, enhances free fatty acid oxidation in
muscles, reduces the accumulation of free fatty acid in liver, and insulin signaling, reduces hepatic
glucose output, and improves insulin sensitivity [41].
IGF-1 and 2 were reported to be equally effective at promoting chick embryonic myoblast
differentiation and fusion [42], whereas IGF-2 was more effective than IGF-1 during turkey
embryonic myoblast proliferation and differentiation. IGFs were found to stimulate in chicken and
turkey MSC proliferation [89, 90], and treatment with IGF-1 and IGF-2 enhanced MSC growth.
When the effect of IGF-1 on chicken myoblast proliferation was examined IGF-1 dose-
dependently increased myoblast numbers. Supplementation with 10, 100, or 1,000 ng/mL of IGF-
1 for 24 h increased numbers by 9.5%, 63.1%, and 66%, respectively. The optimal concentration of
IGF-1 for stimulating proliferation was found to be 100 ng/mL [40].

Myostatin
Myostatin (growth and development factor-8, MSTN) is present in SM, and muscle mass in
MSTN-null mice was reported to be dramatically enhanced due to increased muscle fiber levels
(hyperplasia) and sizes (hypertrophy) [91,92]. Furthermore, the double-muscled phenotype
in cattle has been linked to the MSTN gene. Muscular hypertrophy in Belgian Blue double-
muscled (BBDM) cattle is caused by an 11-base-pare deletion, and this naturally occurring
mutation inactivates the MSTN gene and results in a huge increase in SM mass [93]. Cell culture
experiments have also revealed that MSTN inhibits the proliferation and differentiation of
cultured muscle cell lines and primary myogenic cells [93], which suggests siRNA1 and siRNA5
might reduce MSTN gene expression and boost sheep meat yield [94]. Some natural compounds
[95–97] and inhibitory peptides targeting MSTN [21,98], such as Ac-MIF1 and Ac-MIF2-NH2,
have been reported to inhibit the effect of MSTN significantly and enhance cell proliferation [99].
Additionally, in vivo and in vitro data showed that FMOD prevents muscle aging by decreasing the
activity of MSTN protein [92]. These findings indicate that MSTN gene sequence modification
and MSTN inhibition are good strategies for the mass production of cultured meat from
agricultural animals.

Transforming growth factor β

TGF-β1 is involved in muscle repair through MSC activation, connective tissue development, and
immune response regulation [100]. TGF-β1 also attenuated MSC activation when used with HGF
and when administered during early MSC activation (0–48 h) or the proliferative phase (48–96
h) TGF-β1 maintained or induced MSC quiescence, respectively, based on MyoD protein levels
[101]. TGF-β1 can influence MSC proliferation and myogenesis, enhance mature SM mass, and

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 Ahmad et al.

regulate intramuscular fibrogenesis; it also plays critical roles in muscle formation. TGF-β1 was also
demonstrated to restrict myogenesis in 2D cultures while increasing myogenesis in 3D cultures,
which makes it a useful regulator of in vitro muscle tissue formation [102,103].

ROLE OF HORMONES IN MYOGENESIS FOR CULTURED

MEAT PRODUCTION
Insulin
Insulin is a hormone that facilitates MSC survival and self-renewal in vitro [55]. Insulin maintains
the differentiation power of myoblasts in vitro [104], and in combination with IGF-2, insulin
augments the self-renewal property of MSCs [105], which is necessary for the initiation of culture
meat production. Insulin works by activating its receptor and the PI3K/AKT cascade to promote
cell survival and also activates integrin, which is required for cell survival during niche reformation
after passage. In addition, insulin is needed for human ESC survival on E-cadherin-coated
surfaces and in suspension, which suggests its involvement in cell-to-cell adhesion [55]. Insulin is a
potent mitogen that promotes DNA synthesis and MSC proliferation and has a dose-dependent
stimulatory impact on MSC differentiation and fusion. Insulin appears to promote myogenesis and
differentiation by boosting the production of myogenin and myosin heavy chain (MHC) isoforms,
and insulin administration thickens myotubes, which suggests a role in insulin-induced hypertrophy
[106]. Accordingly, insulin is an important factor for cell survival and growth during the initial stage
of cultured meat.

Testosterone
Testosterone primarily interacts with androgen receptors in SM to promote muscle growth. In SM,
testosterone has a variety of ergogenic, anabolic, and anti-catabolic functions that dose-dependently
contribute to muscle strength and hypertrophy [107,108]. In muscle, testosterone promotes
muscle hypertrophy by stimulating protein synthesis and inhibiting protein degradation, and when
these effects merge, testosterone stimulates muscle hypertrophy. Furthermore, the interaction
between testosterone and intracellular androgen receptor modulates specific physiological signals
[39,108]. Testosterone dose-dependently increases MSC numbers and muscle fiber sizes [61,62].
A significant increase in myoblasts numbers, which generated bigger myofibers, was observed
when MSCs were treated with testosterone. However, the effect of testosterone on myogenesis in
chickens and the mechanism by which testosterone regulates SM development remain unclear. In
addition, testosterone significantly increased the cross-sectional area and density of myofibers in
muscle [62].
Androgens and estrogens also significantly influence muscle physiology and metabolism and
are involved in muscle mass development, maintenance, and repair [109]. Although estrogens and
androgens both have favorable effects on muscle, androgens play a major role in the control of
muscle physiology and also influence muscle mass by decreasing protein breakdown and autophagy
[110]. Overall, estrogens and androgens regulate muscle formation and maintenance.

Glucocorticoids
Glucocorticoids, primarily cortisol, have substantial effects on human SM. Cortisol regulates SM
energy homeostasis and metabolism [111,112]. Glucocorticoid receptors are present in the ECM
of mammalian SM fibers and signal quickly through the mitogen-activated protein kinase pathway
[113,114]. Dexamethasone is a synthetic glucocorticoid known for its anti-inflammatory and
immunosuppressive properties and is used to suppress muscle degeneration and stabilize muscle

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Growth factors and hormones in cultured meat production

strength [115]. At a concentration of 25 nM dexamethasone increased myogenic proliferation

[116], and treatment with dexamethasone has also been reported to increase MSC myogenic
differentiation marked by advanced sarcomere formation [105,116,117]. Also, the addition of
dexamethasone to muscle cell cultures enhances myogenesis.

Thyroid hormones
SM is a primary target of thyroid hormones [118], which are important for stimulating MSC
proliferation and differentiation and muscle recovery and myogenesis [119,120]. Thyroid
hormones enhance the numbers and diameters of muscle fibers and also help determine muscle
fiber distribution patterns [121], but high or low thyroid hormone levels promote muscle atrophy
and impede muscle regeneration [121,122]. Peroxisome proliferator-activated receptor gamma
coactivator 1-alpha (PPARGC-1α) is a major regulator of mitochondrial production and is
significantly up-regulated by thyroid hormones. One study reported that the presence of thyroid
hormones enhanced mitochondrial numbers, protein synthesis, and basal O2 consumption,
adenosine triphosphate (ATP) turnover, and maximum respiratory capacity [118]. SM contains
type 2 and 3 iodothyronine deiodinases (DIO2 and DIO3, respectively). DIO2 is a strictly
controlled enzyme that catalyzes the outer-ring monodeiodination of the secreted prohormone
tetraiodothyronine (T4) to produce active hormone tri-iodothyronine (T3), which can either stay in
myocytes, signal through nuclear receptors, or exit cells and interact with the extracellular pool [121,
123]. Thyroid hormone signaling plays an important role in SM development and regeneration, and
T3 signaling is crucial for SM development [123].

ANIMAL TISSUE CULTURE TECHNIQUES

Cell culture is a process whereby cells are grown in engineered environments, and animal cell
culture is one of the most widely utilized research techniques in the biological sciences. Using tissue
engineering techniques, in vitro meat production provides a unique means of producing meat
without using animals [124]. The basic technology for in vitro meat production involves cultivating
muscle tissue in a bioreactor. Starter cells for meat production can be obtained by live animal biopsy
and then cultured in vitro [124,125]. The development of a basic culture medium has allowed the
growth and differentiation of various cells studied under various conditions [126]. Interestingly,
optimal in vitro culture temperature, pH, CO2, O2, osmolality, and nutrition requirements may
not be identical to in vivo conditions. Furthermore, sterile cultivation conditions, a constant
supply of nutrients, and complex incubation conditions are required [127]. Although nutrients,
adhesion proteins, and GFs are abundant in fetal bovine serum (FBS), the exploration of alternative
media and the creation of serum-free media formulations have attracted global interest due to
shortcomings in in vitro data quality and repeatability and animal welfare issues. The majority of
studies on the topic indicate that chemically specific serum-free mediums are the ultimate goal
[128, 129]. Research efforts continue to be directed toward the development of optimal culture
conditions that improve the efficacy of cultured meat production.
GFs are signaling molecules that stimulate growth, proliferation, and differentiation by
triggering signaling cascades after binding to their receptors. GFs are important cell culture media
components and provide signals for growth, structural stability, and cell-to-cell communication.
However, GFs are expensive, and thus, some alternative means of synthesizing GFs are needed.
Important GFs can be sourced from the genomes of different species. Similar sequences can be
found by searching publicly available genome databases and synthesized or purchased from an
available vendor. Recombinant DNA technology can then be used to insert sequences of interest

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 Ahmad et al.

into bacterial systems. Using this strategy, active GFs, such as FGF-2, IGFs, PDGF-BB, and TGF-
β1 were effectively developed in Escherichia coli, and soluble GFs from cattle, chickens, and fish were
produced [130].

CONCLUSION
GFs and hormones are important media components that regulate MSCs and improve cultured
meat production. After binding to their receptors, these factors deliver the signals for growth,
structural stability, cell-to-cell communication, proliferation, and differentiation. However, they
are expensive media components, and thus, optimization and cheaper ways of synthesizing GFs
and hormones are required. This review describes the importance of the regulatory roles played
by different GFs (FGF-2, IGF-1, PDGF-BB, and TGF-1) and hormones (insulin, testosterone,
glucocorticoids, dexamethasone, and thyroid hormones) during myogenesis and cultured meat
production.

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